The Regulation of Body and Wing Disk Growth in Manduca Sexta
A key question in developmental biology is how organisms attain a final size. Deviations in growth patterns can produce different/new phenotypes and these changes can play fundamental roles in ecology and evolution. The size of an organism and of its constitutive organs is determined by the growth rate and the duration of the growing period. In insects, peptide hormones such as insulin-like growth factors have been shown to be involved in determining the growth rates by coordinating metabolism, cell proliferation and cell size. In contrast, steroid hormones, such as ecdysone, are involved in determining life stage transitions, and thus the termination of the growing period. Although it is clear that insulin and steroid hormones are both involved in the regulation of growth, the ways in which these two regulators interact is yet to be determined. Furthermore, it is not clear how organs and body growth are coordinated during development to arrive to their correct proportions. In this study, using the tobacco hornworm Manduca sexta and its wings as a model system, I examine the developmental mechanisms involved in the regulation of organ growth and how developmental processes can drive morphological evolution. First, I examine how the hormonal events that take place during the termination of the body growth period affect wing disk growth. Second, by using gene expression assays and in vitro cultures, I examine the interaction between bombyxin, the Lepidopteran insulin-like growth factor, and ecdysone, the molting hormone, and their contributions to wing imaginal disk growth. Finally, by using three different size strains of M. sexta, I examine the developmental basis of the allometric relationship between the wings and the body. My results show that during the final instar of M. sexta larval development, wing imaginal disks are sensitive to the hormonal events that terminate the growth period. Furthermore, I show that the bombyxin requirement for wing disk growth is restricted to the early days of the final instar unlike the constitutive effects seen in other species. After the larva has passed a particular critical weight, bombyxin is not necessary for wing disk growth, although its absence does decrease the growth rate. In contrast, ecdysone is required for promoting the growth of wing imaginal disks primarily through its stimulation of cell proliferation. Finally, I show how selection on body size has unpredictable consequence for the response of wing size. These results demonstrate how specific allometries have a developmental basis in the cross-talk of the various signals that regulate growth itself. Therefore, direct selection on allometric relationships may not need to be strong in order to hold scaling relationships constant, at least over short evolutionary periods.
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